Interference at several temporal and spatial scales between two chestnut insects

Detection of interspecific competition between insects is often sensitive to scaling. We give an example of scale-dependent interference between the weevil Curculio elephas and the moth Cydia splendana, which both have larvae that develop in the fruits of chestnut Castanea sativa. Measures at three scales were considered: chestnut, husk (with one to three fertile fruits) and tree. Data come from observations in the field over 14 years, complemented by experiments done directly in trees. Data on individual chestnut fruits revealed a marked statistical interference between the two insects. Experiments demonstrated that presence of a moth larva in a fruit usually inhibits weevil egg-laying. Conversely, weevil presence does not strongly modify moth larval behavior. Cases of double infestation often correspond to fruits first attacked by the weevil. With measures on husks, interference between the two insects was observed only in some trees; its intensity was always weaker than in the chestnuts themselves. At the scale of entire trees, rates of infestation by each insect are not correlated. Interference in chestnut fruits is interpreted by assuming that the weevil female either is sensitive to a repellent molecule originating from a moth larva or its frass, or can detect moth larval sounds. Mechanisms governing infestation rates from data per tree are discussed in relation to those found at fruit scale and to plant-insect interactions. The need to estimate available resources both from quantitative and qualitative points of view is emphasized.     

Infestation rate of argan fruit (Argania spinosa) by the Mediterranean fruit fly (Ceratitis capitata) in relation to phenology and maturation of the fruit

Argan (Argania spinosa (Sapotaceae)) forest covers about 700,000 ha in the Souss region (Morocco). The medfly Ceratitis capitata Wiedemann (Diptera: Tephritidae) is known to develop in argans. In this study, we examine variations in medfly infestation rates according to argan phenology from 1987 to 1990. Argan phenology varies greatly among trees, total fruit drop spreading from late March to mid-July; moreover, an exceptional fructification may occur in Autumn in some trees. From February to about May, the infestation rate is low except in fruit produced during autumnal fructification. In June and July, the rate increases greatly reaching 100% in some trees. Ripe and ripening argans are the most heavily infested; only 10% of immature fruits on the average are attacked by the medfly. Fallen argans are more heavily infested than fruit still on the tree. Productivity per argan averaged from one to three adult medflies. Our results demonstrate that argan forest in the Souss acts as an enormous reservoir of medflies for the nearby citrus orchards.

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Résumé L'arganeraie (Argania spinosa) couvre environ 700000 ha dans la région du Souss, au Maroc. La mouche méditerranéenne des fruits, Ceratitis capitata, est connue pour se développer dans les arganes. Des études ont été menées, de 1987 à 1990, pour étudier comment le taux d'infestation des arganes variait selon leur degré de maturation. La phénologie des arganiers varie fortement selon les arbres: par exemple la chute totale des fruits s'étale de fin mars à mi-juillet. De plus, une fructification peut être observée en automne mais elle ne concerne qu'un nombre limité d'arbres. De février à mai environ, le taux d'infestation des arganes reste faible, sauf pour les fruits produits pendant la fructification de l'automne précédent. En juin en juillet, le taux augmente fortement atteignant 100% dans certains arbres. Les arganes mûres ou mûrissantes sont les plus attaquées; seuls 10% en moyenne des fruits immatures sont infestés. Les fruits tombés sont toujours (sauf à la fin) nettement plus attaqués que ceux encore présents dans l'arganier. De une à trois Cératites adultes sont formées à partir d'une argane. Nos résultats démontrent que la forêt d'argane du Souss constitue un réservoir gigantesque de la Cératite pour les infestations des vergers d'agrumes proches.

     

The impact of the discovery of vertebrate-type steroids and peptide hormone-like substances in insects

L'influence de quelques variables liées aux plantes de Phaseolus vulgaris (date de maturation, hauteur et nombre de graines dans les gousses) sur la ponte de la bruche du haricot Acanthoscelides obtectus a été étudiée dans la nature sur 20 pieds de haricot dont toutes les gousses ont été suivies puis récoltées.Les résultats montrent une bonne corrélation entre le nombre d'oeufs émis dans une gousse et le nombre de jours avec présence d'adultes sur cette gousse. La date de maturation influence fortement la contamination des gousses. Au niveau spatial, les pieds situés dans des zones non désherbées, où peuvent s'alimenter les adultes, sont les plus contaminés. Les autres variables semblent avoir moins d'importance.On note une très forte agrégativité des pontes dans certaines gousses qui sont, en général, parmi les premières mûres, mais cette agrégativité persiste même si les femelles ont le choix, sur un même pied, entre plusieurs gousses arrivées ensemble à maturité.     

Coin-flipping plasticity and prolonged diapause in insects: example of the chestnut weevil Curculio elephas (Coleoptera: Curculionidae)

Spreading of emergence over several years due to prolonged diapause in some larvae was shown in the chestnut weevil. Depending on the year the larvae buried themselves in the ground, 32–56% of live adults emerged after 2 or 3 years of underground life. Variability in the duration of diapause was assumed to correspond to tactics of adaptative coin-flipping plasticity. This plasticity must allow the chestnut weevil to respond to the unpredictability of its habitat as measured by the irregularity of chestnut production and summer drought. Indeed, fecundity and adult longevity did not lessen after 2 years of underground life. No drastic decrease in the population size of weevils occurs after bad years; for instance when the number of chestnuts on the study tree is less than 10 000, passers-by can collect all the fruit and about 95% of larvae developing in chestnuts are destroyed. Diapause nature (simple or prolonged) may be related to moisture and gas rates in the ground from October to December. These factors acting in autumn are not known to be involved in the physiological mechanisms that control the production of chestnuts.     

Oviposition pattern of phytophagous insects: on the importance of host population heterogeneity

Frequency distributions of insect immatures per host are often fitted to contagious distributions, such as the negative binomial, to deduce oviposition pattern. However, different mechanisms can be involved for each theoretical distribution and additional biological information is needed to correctly interpret the fits. We chose the chestnut weevil Curculio elephas, a pest of the European chestnut Castanea sativa, as a model to illustrate the difficulties of inferring oviposition pattern from fits to theoretical distributions and from the variance/mean ratio. From field studies over 13–16 years, we show that 20 out of the 31 yearly distributions available fit a negative binomial and 25 a zero-inflated Poisson (ZIP). No distribution fits a Poisson distribution. The ZIP distribution assumes heterogeneity within the fruit population. There are two categories of host: the first comprises chestnuts unsuitable for weevil oviposition or in excess relative to the number of weevil females, and the second comprises suitable fruits in which oviposition behavior is random. Our results confirm this host heterogeneity. According to the ZIP distribution, the first category of hosts includes on average 74% of the chestnuts. A negative binomial distribution may be generated by either true or false contagion. We show that neither interference between weevil females, nor spatial variation in the infestation rate exist. Consequently, the observed distributions of immatures are not the result of false contagion. Nevertheless, we cannot totally exlude true contagion of immatures. In this paper we discuss the difficulty of testing true contagion in natural conditions. These results show that we cannot systematically conclude in favour of contagion when fitting a distribution such as the negative binomial or when a variance/mean ratio is higher than unity. Received: 22 September 1997 / Accepted: 15 December 1997     

Geostatistical analysis of the distribution of NH(4)(+) and NO(2)(-)-oxidizing bacteria and serotypes at the millimeter scale along a soil transect

Soil is known to be heterogeneous for different activities at several spatial scales. Most studies have focused on macro- and meso-scales but micro-scales are rarely addressed. Hence, the spatial structure of NH(4)(+)- and NO(2)(-)-oxidizers and of various serotypes of the latter was studied along two transects of approximately 10 cm, with two micro-samples taken from each millimeter. The presence of NH(4)(+)- and NO(2)(-)-oxidizers in a micro-sample was detected using colorimetric tests for the presence or absence of NO(2)(-) in cultures of the micro-samples. Geostatistics was used to determine the range of spatial influence of the bacterial types. For both types, semi-variograms indicated a non-random spatial pattern. The spatial dependence ranged from 2 to 4 mm for NO(2)(-)- and NH(4)(+)-oxidizers respectively, and the two bacterial types were not independently spatially located. Among the six serotypes of NO(2)(-)-oxidizers, only one exhibited a spatial dependence. The existence of a spatial structure at the millimeter scale suggests that micro-scale sampling should be employed for soil studies. Therby, data on bacterial populations and activities can be referred to a spatial scale which is meaningful to these organisms.     

Clutch size manipulations in the chestnut weevil, Curculio elephas: fitness of oviposition strategies

We test the adaptive value of clutch size observed in a natural population of the chestnut weevil Curculio elephas. Clutch size is defined as the number of immatures per infested chestnut. In natural conditions, clutch size averages 1.7 eggs. By manipulating clutch size in the field, we demonstrate that deviations from the theoretical ”Lack clutch size”, estimated as eight immatures, are mainly due to proximate and delayed effects of clutch size on offspring performance. We show the existence of a trade-off between clutch size and larval weight. The latter, a key life-history trait, is highly correlated with fitness because it is a strong determinant of larval survival and potential fecundity of offspring females. The fitness of different potential oviposition strategies characterized by their clutch sizes, ranging from one to nine immatures, was calculated from field- estimated parameters. Chestnut weevil females obtain an evolutionary advantage by laying their eggs singly, since, for instance, fitness of single-egg clutches exceeds fitness of two-egg clutches and four-egg clutches by 8.0% and 15.1% respectively. Received 4 August 1999 / Accepted: 7 October 1999